EXTANT SEED PLANTS

Plant woody, evergreen; nicotinic acid metabolised to trigonelline, (cyanogenesis via tyrosine pathway); primary cell walls rich in xyloglucans and/or glucomannans, 25-30% pectin [Type I walls]; lignins rich in guaiacyl units; true roots present, apex multicellular, xylem exarch, branching endogenous; arbuscular mycorrhizae +; shoot apical meristem multicellular, interface specific plasmodesmatal network; stem with ectophloic eustele, endodermis 0, xylem endarch, branching exogenous; vascular tissue in t.s. discontinuous by interfascicular regions; vascular cambium + [xylem ("wood") differentiating internally, phloem externally]; wood homoxylous, tracheids +; tracheid/tracheid pits circular, bordered; sieve tube/cell plastids with starch grains; phloem fibers +; stem cork cambium superficial, root cork cambium deep seated; nodes ?; stomata ?; leaf vascular bundles collateral; leaves spiral, simple, axillary buds?, prophylls [including bracteoles] two, lateral, veins -5 mm/mm² [mean for all non-angiosperms 1.8]; plant heterosporous, sporangia eusporangiate, on sporophylls, sporophylls aggregated in indeterminate cones/strobili; true pollen [microspores, i.e. no distal pore for release of gametes] +, grains mono[ana]sulcate, exine and intine homogeneous, ovules unitegmic, crassinucellate, megaspore tetrad tetrahedral, only one megaspore develops, megasporangium indehiscent; male gametophyte development first endo- then exosporic, tube developing from distal end of grain, to ca 2 mm from receptive surface to egg, gametes two, with cell walls, with many flagellae; female gametophyte endosporic, initially syncytial, walls then surrounding individual nuclei; seeds "large", first cell wall of zygote transverse, embryo straight, endoscopic [suspensor +], short-minute, with morphological dormancy, white, cotyledons 2; plastid transmission maternal; two copies of LEAFY gene, PHY gene duplication [N/O//A/C and P//BE lines], mitochondrial nad1 intron 2 and coxIIi3 intron present.

MAGNOLIOPHYTA

Plant woody, evergreen; lignans, O-methyl flavonols, dihydroflavonols, triterpenoid oleanane, non-hydrolysable tannins, quercetin and/or kaempferol +, apigenin and/or luteolin scattered, [cyanogenesis in ANITA grade?], lignins derived from both coniferyl and sinapyl alcohols, containing syringaldehyde [in positive Maüle reaction, syringyl:guaiacyl ratio less than 2-2.5:1], and hemicelluloses as xyloglucans; root apical meristem intermediate-open; root vascular tissue oligarch [di- to pentarch], lateral roots arise opposite or immediately to the side of [when diarch] xylem poles; origin of epidermis with no clear pattern [probably from inner layer of root cap], trichoblasts [differentiated root hair-forming cells] 0; stem with 2-layered tunica-corpus construction; wood fibers and wood parenchyma +; reaction wood ?, with gelatinous fibres; starch grains simple; primary cell wall mostly with pectic polysaccharides; tracheids +; sieve tubes eunucleate, with a sieve plate and cytoplasm with P-proteins, companion cells from same mother cell that gave rise to the sieve tube; nodes unilacunar [1:?]; stomata with ends of guard cells level with pore, paracytic, outer stomatal ledges producing vestibule; leaves with petiole and lamina [the latter formed from the primordial leaf apex], development of venation acropetal, 2ndary veins pinnate, fine venation reticulate, veins (1.7-)4.1(-5.7) mm/mm², endings free; flowers perfect, polysymmetric, parts spiral [esp. the A], free, development in general centripetal, numbers unstable; P not sharply differentiated, outer members not enclosing the rest of the bud, smaller than inner members; A many, with a single trace, introrse, filaments stout, anther ± embedded in the filament, tetrasporangiate, dithecal, with at least outer secondary parietal cells dividing, each theca dehiscing longitudinally by action of hypodermal endothecium, endothecial cells elongated at right angles to long axis of anther; tapetum glandular, binucleate; microspore mother cells in a block, microsporogenesis successive, walls developing by centripetal furrowing; pollen subspherical, binucleate at dispersal, trinucleate eventually, tectum continuous or microperforate, ektexine columellar, endexine thin, compact, lamellate only in the apertural regions; nectary 0; G free, several, ascidiate, with postgenital occlusion by secretion, few [?1] ovules/carpel, ovules marginal, anatropous, bitegmic, [outer integument often largely subdermal in origin, inner integument dermal], micropyle endostomal, integuments 2-3 cells thick, megasporocyte single, megaspore lacking sporopollenin and cuticle, chalazal, female gametophyte four-celled [one-modular, nucleus of egg cell sister to one of the polar nuclei], stylulus short, hollow, stigma ± decurrent, dry [not secretory]; P deciduous in fruit; seed exotestal; pollen germinating in less than 3 hours, tube elongated, growing at 80-600 µm/hour, with callose plugs and callose-based walls, penetrating between cells, siphonogamy, penetration of ovules within ca 18 hours, distance to first ovule 1.1.-2.1 mm; double fertilisation +, endosperm diploid, cellular [first division oblique, micropylar end initially with a single large cell, chalazal end more actively dividing], copious, oily and/or proteinaceous, embryo cellular ab initio, minute; germination hypogeal, seedlings/young plants sympodial; Arabidopsis-type telomeres [(TTTAGGG)_n]; whole genome duplication, single copy of LEAFY and RPB2 gene, knox genes extensively duplicated [A1-A4], AP1/FUL gene, paleo AP3 and PI genes [paralogous B-class genes] +, with "DEAER" motif, SEP3/LOFSEP and PHYA + C/PHYB + E gene pairs.

Evolution. Possible apomorphies for flowering plants are in bold. Note that the actual level to which many of these features, particularly the more cryptic ones, should be assigned is unclear, because some taxa basal to the [magnoliid + monocot + eudicot] group have been surprisingly little studied, there is considerable variation between families in particular for several of these characters, and also because details of relationships among gymnosperms will affect the level at which some of these characters are pegged. For example, if reticulate-perforate pollen is optimized to the next node on the tree (see Friis et al. 2009 for a discussion), it effectively makes the pollen morphology of the common ancestor of all angiosperms ambiguous....

NYMPHAEALES [AUSTROBAILEYALES [[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]]]]: vessels +, elements with scalariform perforation plates, axial parenchyma diffuse or diffuse-in-aggregate; tectum reticulate-perforate [here?]; ?genome duplication; "DEAER" motif in AP3 and PI genes lost, gaps in these genes.

ERICALES [ASTERID I + II]: ovules tenuinucellate.

ASTERID I + II: ellagic acid 0, proanthocyanidins not common; inflorescence cymose; C forming a distinct tube; A epipetalous, = and opposite sepals or P [polyandry (secondary) very uncommon indeed].

ASTERID I: G [2]; loss of introns 18-23 in d copy of RPB2 gene.

BORAGINACEAE + VAHLIACEAE + GENTIANALES + LAMIALES + SOLANALES: (8-ring deoxyflavonols +); vessel elements with simple perforation plates; C tube initiation late [sampling!]; [vascularised] nectary at base of G; style long.

LAMIALES + SOLANALES: iridoids, myricetin, non-hydrolysable tannins usu. 0; nodes 1:1; K connate, anther sacs with placentoids.

SOLANALES Dumortier  Main Tree, Synapomorphies.

O-methyl flavonols (flavones) +; pollen tube usu. with callose; K persistent in fruit; endosperm development? - 5 families, 165 genera, 4080 species.

Evolution. Stem group Solanales may date from the Campanian-Santonian 86-82 million years before present, diversifying 78-76 million years before present (Wikström et al. 2001: Sphenoclea not included); Janssens et al. (2009) date stem group Solanales to 101±11.8 million years ago; Bremer et al. (2004) date the stem clade to ca 106 million years ago and the crown clade to ca 100 million years.

Phylogeny. Montiniaceae are placed next to Solanaceae + Convolvulaceae (B. Bremer 1996, see also Soltis & Soltis 1997; cf. Takhtajan 1997 - in Hydrangeales). D. Soltis et al. (2000) found strong support for the association of Montinia and Hydrolea; Sphenoclea was not included. With the inclusion of the latter and broader sampling (three genera) in Montiniaceae, B. Bremer et al. (2002) found strong support for the association of Sphenoclea and Hydrolea, but only just above 50% for the association of Montiniaceae with that pair. The topology of the backbone of the tree here follows that of the latter paper.

Includes Convolvulaceae, Hydroleaceae, Montiniaceae, Solanaceae, Sphenocleaceae.

Synonymy: Cestrales Schlechtendahl, Convolvulales Dumortier, Cuscutales Bartling, Hydroleales R. Brown, Nolanales Lindley, Sphenocleales Doweld - Solananae Reveal - Convolvulopsida Brongniart, Solanopsida Brogniart

Montiniaceae [Sphenocleaceae + Hydroleaceae]: alkaloids/; petiole bundle(s) arcuate; inflorescence terminal.

MONTINIACEAE Nakai   Back to Solanales

Shrubs, trees (lianes); plants with a peppery smell; route II secoiridoids +, plant slightly tanniniferous; cambium storied or not; pits vestured; young stem with a vascular cylinder (separate bundles; medullary bundles +); nodes 1:1(?-11:11 - Kaliphora); crystal sand, acicular crystals and styloids usu. all +; petiole arc of (rounded) bundles (+ additional strands); tuft of hairs at nodes; (stomata anisocytic - some Grevea); leaves also opposite; bracteoles 0; flowers imperfect, small, K small, C free [absolutely so - Montinia], valvate or not, nectary vascularised; staminate flowers: 3-4(-5)-merous, anthers extrorse, anthers ± basifixed, pollen grains large, pistillode minute; carpellate flowers: 4-merous; A staminodial (0), ovary inferior, placentation intrusive parietal-subaxile, 1-12 (campylotropous - Kaliphora) ovules/carpel, nucellus and endothelium?, style short, stout, hollow, stigma with 2 large lobes; fruit a capsule; seeds winged, exotesta lignified, periclinal walls thickened, adjacent wall of mesotesta also thickened [Montinia]; fruit indehiscent, placentae at least initially fleshy; testa thin-walled, ± pulpy when wetted [Grevea]; fruit a drupe; (exotesta not persistent Grevea); endosperm +/0, ?development, hemicellulosic, walls layered, cotyledons foliaceous, radicle oblique; cotyledonary petioles connate [Montinia]; n = 16 [Kaliphora], 34 [Montinia].

3[list]/5. Africa and Madagascar (map: from Milne-Redhead & Metcalfe 1955; Verdcourt 1975; Bosser 1990; Brummit 2007 [C. and W. Africa]). [Photos - Kaliphora, Montinia Fruits © Serban Procheŝ.]

• Montiniaceae are woody plants that can be recognised by the persistent short style with large stigmatic lobes that cap the smooth fruit; the ovary is inferior. The often 4-merous flowers are small, with free, valvate petals and extrorse, more or less basifixed anthers. Crushed tissue may have a spicy or acrid smell.

Chemistry, Morphology, etc. Pericyclic fibers may be poorly developed; Grevea has vascular bundles in the pith. The axillary tufts of hairs are at best poorly developed in Kaliphora and are probably not apomorphic for the family. The pollen (Hideux & Ferguson 1976) is rather like that of some Araliaceae.

See Milne-Redhead and Metcalfe (1955) for general morphology and anatomy, Hegnauer (1973, 1990, as Saxifragaceae) for chemistry, Dahlgren et al. (1977) for germination and iridoids, Ramamonjiarisoa (1980), Carlquist (1989), and Gregory (1998) for vegetative anatomy, Krach (1976, 1977) and Takhtajan and Trifonova (1999) for testa anatomy, and Ronse Decraene (1992) and Ronse Decraene et al. (2000a) for floral details.

Previous Relationships. Montiniaceae have been hard to place, and have generally been included somewhere around Saxifragaceae. Cronquist (1981) included them in his heterogeneous Grossulariaceae, while Takhtajan (1997) placed them as a separate family (Kaliphoraceae were adjacent) in his Hydrangeales.

Synonymy: Kaliphoraceae Takhtajan

Sphenocleaceae + Hydroleaceae: placentae swollen [?level]; endosperm cellular, at most scanty, with multicellular micropylar and chalazal haustoria.

SPHENOCLEACEAE Baskerville   Back to Solanales

Rather fleshy annual herbs; fructose with isokestose linkages, cyclic thiosulphinates [zeylanoxides] +; cork ?mid-cortical; cortical air spaces +; stomata tetracytic; inflorescences spicate; C tube formation early, C imbricate, pollen trinucleate, nectary 0; G ± inferior, many ovules/carpel, style short, stigma capitate; fruit capsular, circumscissile; exotestal cells with inner walls thickened; n = 12, 16, 20, etc.

1/2. Pantropical. [Photo - Habit © B. Hammel]

• Sphenocleaceae are erect fleshy annuals of damp or flooded habitats that have dense spikes of small flowers with more or less inferior ovaries. The small fruits are circumscissile, the lid coming off just below the point of origin of the calyx. Older stems have a green reticulation marking the plates of cells in the tissue below bounding the cortical air spaces.

Chemistry, Morphology, etc. Corolla tube formation is of the early type, and the corolla lobes are characteristically incurved; the lateral veins of adjacent lobes are fused producing commissural veins. Some information is taken from Monod (1980).

Previous Relationships. Sphenocleaceae, along with Hydrolea, another genus of uncertain position, were placed near Boraginaceae by Cosner et al. (1994). However, in morphological studies (e.g. Gustafsson & Bremer 1995) Sphenocleaceae seem to be satisfactorily positioned well within Asterales. Indeed, Sphenocleaceae have often been associated with Campanulaceae (e.g. they are placed in Campanulales by Takhtajan 1997), although they lack latex.

HYDROLEACEAE Berchtold & J. Presl   Back to Solanales

Herb to shrubby; chemistry?; axillary-sublateral thorns or not; cork?; vessel elements?; stomata?; leaf margin toothed to entire; flowers 4-5-merous, K basally connate, C tube formation late; A versatile, nectary 0/+; G [2(-4)], diagonal, placentae bilobed, many mostly pleurotropous ovules/carpel, antipodals degenerating early, funicular bundle absent, styles +, stigma slightly funneliform or capitate; fruit a septi- (and loculi-)cidal capsule (irregularly dehiscent); seeds longitudinally ridged and ruminate, exotestal cells thin-walled, endotestal cells tanniniferous, with a cuticle; n = (9) 10 (12).

1/12. Tropical, warm temperate (map: from Davenport 1988; FloraBase 2007). [Photo - Hydrolea Flower © B. Kenney]

• Hydroleaceae are more or less shrubby plants growing near or even in water. They have rather small leaves.

Chemistry, Morphology, etc. Hydrolea appears to lack mycorrhizae. The axillary inflorescences may be cymose. Davenport (1988) suggested that the disc is absent. The two carpels are shown as being oblique by Schnizlein (1843-1870: fam. 147), and this is confirmed by Erbar et al. (2005), even for Hydrolea palustris, which has flowers with the median sepal abaxial. Di Fulvio (1997) notes that the four ventral bundles of the two carpels are all connate in the center of the ovary - cf. Hydrophyllaceae s. str., where there are two or four such bundles. There are no nuclear inclusions.

Details of embryology are taken from Svensson (1925) and di Fulvio (1989b, 1990); Davenport (1988) monographed the genus.

Previous Relationships. Hydrolea has usually been included in Hydrophyllaceae (e.g. Cronquist 1981; Takhtajan 1997). Not only molecular differences but also axile versus parietal placentation and embryological differences (see di Fulvio de Basso 1990) separate the two.

Convolvulaceae + Solanaceae: coumarins, caffeic acid esters, tropane [polyhydroxynortropanes], pyrrolidine, etc. alkaloids, flavonol and flavone glycosides, acylated anthocyanins +, tannins 0; internal phloem +; leaves with conduplicate ptyxis; late C-tube formation, C usu. contorted or induplicate-valvate; G opposite petals, many ovules/carpel, (integumentary tapetum +); K persistent in fruit; young seeds starchy.

Evolution. This clade may have diverged 78-76 million years before present, diversifying perhaps 66-65 million years before present (Wikström et al. 2001). Chrysomelidae-Cassidinae+Hispinae and -Criocerinae beetle larve like members of this clade, especially Convolvulaceae (Schmitt 1988; Jolivet 1988; Buzzi 1994; Vencl & Morton 1999).

Chemistry, Morphology, etc. Pyrrolizidine, tropane and pyrrolidine alkaloids are all synthesised from an ornithine precursor (Hegnauer 1973; Dahlgren 1988).

It is possible that flowers with oblique symmetry may be an apomorphy for this clade.

CONVOLVULACEAE Jussieu, nom. cons.   Back to Solanales

Plant laticiferous; stomata usu. paracytic; leaf margins entire; K quincuncial, large, free; placentoid 0, ovules apotropous, stigma dry.

57[List]/1625 - two groups below. World wide (map: from Meusel et al. 1978; Staples & Brummit 2007).

1. Humbertioideae Roberty

Large tree; chemistry?; vascular bundles collateral; petiole bundle annular; usu. articulated latex canals, or latex cells in the flowers alone; flowers single, axillary, strongly obliquely monosymmetric; A adnate to base of C, filaments bent in bud, style clavate; fruit a few-seeded drupe; endosperm copious; n = ?

1/1: Humbertia madagascariensis. Madagascar.

Synonymy: Humbertiaceae Pichon, nom. cons.

2. Convolvuloideae Burnett

Herbaceous right-twining vines (lianes to 30 m; trees); (ergoline alkaloids + - from clavicipitalean fungi); (cork pericyclic); secondary thickening anomalous; (fibers or sclereids +); unicellular T-shaped hairs common (hairs stellate); leaves (compound; margins lobed, toothed [dentate - Hyalocystis]), 2ndary veins pinnate to palmate; inflorescence usu. a dichasium; tapetum cells multinucleate, pollen pantoporate or 6-polycolpate; G [2(-5)], (false septum - Mina), (1-)2(-4) erect weakly crassinucellate ovules/carpel, placental obturator common, stigmas capitate and with multicellular papillae or punctate and smooth; fruit usu. a variously dehiscent capsule; testa with unbranched bundle, exotesta with papillae or hairs, usu. little thickened, outer hypodermis of small cells, little thickened, inner hypodermis elongated or not, 1+ palisade layers, thickened, 4-8 layers of sclereidal cells underneath, (cells little elongated and thickened - Erycibeae, Maripeae); endosperm nuclear, suspensor endosperm haustorium +, embryo green, curved, cotyledons often folded or coiled, bifid; n = 7-15+; chloroplast gene atpB with 6-15 bp deletion, ycf15 absent, trnF with 150 bp deletion, rpl2 intron 0; sporophytic incompatibility system present.

56/1600: Ipomoea (500 - I. batatas, the sweet potato), Cuscuta (145), Convolvulus (100), Argyreia (90), Jacquemontia (90), Erycibe (75), Merremia (70). World-wide. [Photo - Flower, Fruit.]

Synonymy: Cressaceae Rafinesque, Cuscutaceae Berchtold & J. Presl, nom. cons., Dichondraceae Dumortier, nom. cons., Erycibaceae Meisner, Poranaceae J. Agardh

• Convolvulaceae are recognisable by their usually viny habit and leaves with palmate venation; the plants are often obviously laticiferous. The quincuncial calyx is often notably persistent; the sepals are free, and are rarely green in fruit, often being brown and woody - as in wood roses (Merremia tuberosa) - to scarious. The large, radially symmetric corolla has contorted or induplicate-valvate aestivation and the five stamens are adnate to the tube. The gynoecium is bicarpellate, each loculus usually having two ovules (you rarely need to check this since the calyx is so distinctive), and the seeds have a notably hard, shiny testa.

Evolution. Convolvulaceae are described as being the only asterid family that has seeds showing physical dormancy. This is caused by the thick, hard seed coat found in most members of the family, water initially penetrating the seed only at particular places in the coat (Jayasuriya et al. 2009); indeed, such thick seed coats are decidedly uncommon in the asterids.

The ergoline alkaloids of a number of Convolvulaceae (Ipomoea, Turbina) appear to be synthesized by clavicipitalean fungi, being found in tissues where those fungi also occur (Ahimsa-Müller et al. 2007; Markert et al. 2008).

Cuscuta is a morphologically distinctive parasite - it lacks internal phloem; the corolla is imbricate, with a fimbriate corona especially well developed below and adaxial to the stamens, the ovules are (incompletely) tenuinucellate, the styles separate to connate, and the stigma is capitate to elongated. Its testa anatomy is like that of other members of the family, but the embryo is spirally coiled and (almost) acotyledonous; there is no root. Interestingly, Cuscuta exaltata, at least, has retained most of the genes associated with photosynthesis, perhaps because of their involvement in lipid synthesis (McNeal et al. 2007). For a model of nutrient flow between host and parasite, see Hibberd and Jaeschke (2001).

The flowers often last for only a single day.

Chemistry, Morphology, etc. Glycine betaines are rather commonly accumulated in Convolvulaceae (Rhodes & Hanson 1993), perhaps surprising since it is not a family of halophytes. Wood fluorescence occurs, but not often. Humbertia has hard wood with the odor of sandalwood. The bracts may be adnate to the pedicel and accrescent (wind dispersal: Neuropeltis), or the bracteoles may be much enlarged (e.g. Calystegia). Some Convolvuloideae have flowers with slight oblique disymmetry (Lefort 1951), while the zygomorphy of the flowers of Humbertia is largely positional, indeed, they are drawn as being polysymmetrical by Pichon (1947). The sepals of Humbertia have five traces, but in Convolvuloideae there are fewer; secretory cells are apparently restricted to the flower (Deroin 1993). The corolla tube of some Cuscuta and some other members of the family is strictly speaking a corolla-stamen tube, both contributing integrally to the tubular structure (Prenner et al. 2002). Weberling (1989) described the ovary as being fundamentally gynobasic, although with an apical septum. There are a few, mostly old records of protein crystalloids in the nucleus (Speta 1977; Thaler 1966).

For Cuscuta, see Kuijt (1969) and Heide-Jørgensen (2008); Sherman et al. (2008) describe germination. For chemistry, see Hegnauer (1964, 1989), for seed reserves, see G. Dahlgren (1991), for the rpl2 intron, Downie et al. (1991) and Stefanovic et al. (2002), for a morphological phylogeny, see Austin (1998), for ovary morphology, see Deroin (1999b), for the chloroplast ycf15 gene, see McNeal et al. (2007),for general information, see Staples and Brummitt (2007), for successive cambia, etc., see Rajput et al. (2008). Some other information about Humbertia is taken from Pichon (1947) and K. Kubitzki and H. Manitz (pers. comm.), but the genus is poorly known, especially embryologically.

Phylogeny. Within Convolvuloideae, part of Poraneae (Cardiochlamyeae: Porana itself is polyphyletic), Erycibeae s. str., and a clade made up of all other Convolvuloideae form a basal trichotomy. Cardiochlamyeae have foliaceous bracts and fruits that are utriculate, i.e. they are one-seeded and have a papery pericarp. Erycibe itself (= Erycibeae) has broad, radiate, sessile stigmas and a berry-like fruit; it can look very unlike other members of the family and herbarium specimens are often misidentified. Ipomoea, Convolvulus, and their relatives form an embedded clade that is sister to a rather unexpected clade made up of Poraneae, Cresseae, Dichondreae (with gynobasic styles), some Erycibeae (Maripeae), etc., as well as Jacquemontia. Several members of this latter clade have separate styles or only an at most short common style and long branches (but not Jacquemontia, etc., which have a long common style), and leaf blades with more or less pinnate venation; Jacquemontia could be sister to the other taxa. Despite the sequencing of over 6800 bp, the position of Cuscuteae remains unclear (Stefanovic & Olmstead 2001, 2004; Stefanovic et al. 2002 and esp. 2003); species limits in Cuscuta are difficult (Costea & Stefanovic 2009).

Ipomoea is paraphyletic, within it there is a well-supported spiny pollen clade that comprises some 50% of the larger Ipomoea clade (Manos et al. 2001a).

Classification. For tribal groupings, see Staples and Brummitt (2007); for Cuscuta, see the Parasitic Plants website (Nickrent 1998 onwards) and the Digital Atlas of Cuscuta (Costea 2007 onwards).

SOLANACEAE Jussieu, nom. cons.   Back to Solanales

Herbs to shrubs, branching sympodial; hygroline alkaloids, (withanolides [steroidal lactones]), oligosaccharides, (myricetin) +; roots diarch [lateral roots 4-ranked]; (hairs branched/stellate); wood commonly fluoresces; pits vestured; crystal sand +, esp. in stem; cystoliths +; stomata various; leaves simple to compound; (branching in inflorescence/leaf insertion apparently deviating from normal); (flowers 4 merous; obliquely monosymmetric), anthers often porose, or ± connate and pollen exiting communal apical hole, tapetum cells often 4-nucleate; G [(-5)], oblique, often pseudo-4-locular, usu. many often campylotropous ovules/carpel, chalazal embryo sac haustorium +, stigma capitate or peltate, wet; fruit a berry or septicidal capsule (drupe), K accrescent or not; exotestal walls thickened, endotesta persistent, walls ± lignified; endosperm (helobial, nuclear) +, cotyledons and radicle same width; n = 7-14, etc., chromosomes 1-3 µm long, protein bodies in nuclei.

102[list]/2460 - 8 clades below, but treatment needs work. World-wide, but overwhelmingly tropical America (map: from van Steenis & van Balgooy 1966; Meusel et al. 1978; van Balgooy 1984; Heywood 2007). [Photos - Iochroma Flower, Przewalkskia Fruiting Calyx, Schizanthus Flower.]

1. Schizanthoideae Hunziker

Annual herbs; tropane alkaloids +; cork pericyclic; pericycle fibers 0; flowers strongly monosymmetric, abaxial pair of C connate, forming a keel; A 2 [abaxial-lateral], staminodes 3; endosperm nuclear, embryo curved; n = 10.

1/12. Chile.

The flower is described as having oblique rather than inverted symmetry (Cocucci 1989b), and anther dehiscence is explosive (Cocucci 1989a); for floral evolution, see Pérez et al. (2006: midpoint rooting). Schizanthoideae have distinctive tropane alkaloids, hairs, and pollen (Hunziker 2001).

2. Goetzeoideae Thorne & Reveal

Trees to shrubs; fruit often a drupe; pollen tricolpate, exine echinate, tectum perforate; endosperm at most slight, cotyledons large, fleshy [Goetzea, etc.]; n = 12, 13.

6/8. Most Greater Antilles, but not Jamaica, east Brazil, Madagascar (Tsoala). [Photo - Flower.]

Synonymy: Goetzeaceae Airy Shaw

3. Duckeodendron

Tree; wood with large, open, radial canals [cf. Apocynaceae s. str.]; 1 ovule/carpel; fruit a one-seeded drupe; endosperm slight, embryo U-shaped, cotyledons small.

1/1: Duckeodendron cestroides. Amazonian Brazil.

Synonymy: Duckeodendraceae Kuhlmann

4. Browallioideae Kosteletzky

(Steroid alkaloids - Cestrum); cork superficial or deep-seated; bordered pits +; pericyclic fibers +; A 4 or 5, often didynamous, staminode +/0; n = (7-)11(-13).

10/210: Cestrum (175). South and Central (and North) America.

Synonymy: Cestraceae Schlechtendal, Salpiglossidaceae Hutchinson

5. Schwenckioideae

Annual herbs; pericycle fibers +; flowers monosymmetric, C lobes 3-lobed [Schwenckia and Melananthus]; A 4, didynamous, or 2 + 2 or 3 staminodes; embryo straight, short; n = 10, 12.

4/31. South America.

Petunioideae [Nicotianoideae + Solanoideae] (if this clade exists): (tropane alkaloids [calystegines] +).

6. Petunioideae

Cork superficial (deep-seated); bordered pits +; pericyclic fibers +(0); druses 0(+); (flowers monosymmetric); A 4(-5), usu. of two lengths; embryo also slightly curved; n = 7-11.

13/160: Brunfelsia (45). Central and South America.

Nicotianoideae + Solanoideae: (nicotine [pyridine alkaloid] +); (cotyledons accumbent); x = 12.

7. Nicotianoideae Miers

Cork superficial; pericyclic fibers +/0; A 4 (staminode +), 5, (of two lengths); embryo straight (curved), radicle short; n = (7-11).

8/125: Nicotiana (95), Petunia (35). Mostly Australian, also North and South America, Africa.

Synonymy: Nicotianaceae Martynov

6. Solanoideae Kosteletzky

(Small tree); (steroidal alkaloids +); pits not vestured; (crystal sand +) [level?]; A 5 (4), (of different lengths), base of the filament [stapet] often enlarged, with lobes, etc., (style gynobasic; G 3, or subdivision of carpels into 1-seeded units - "Nolanaceae"); fruit a berry (drupe; circumscissile capsule - Hyoscyameae; schizocarp; K highly accrescent); seeds flattened; endosperm cellular, embryo curved, often coiled; (n = 10-15).

62/1940: Solanum (1250-1700), Lycianthes (200), Lycium (90), Nolana (90), Physalis (80). World-wide, but esp. South America and others N. temperate. [Photo - Flower.]

Synonymy: Atropaceae Martynov, Daturaceae Rafinesque, Hyoscyamaceae Vest, Lyciaceae Rafinesque, Nolanaceae Dumortier, nom. cons., Sclerophylacaceae Miers

• Solanaceae are quite often rather foetid herbs or weak shrubs with polysymmetric or weakly, rarely strongly, monosymmetric sympetalous flowers - when monosymmetrical, the flowers are held so that the odd or median petal is in the adaxial position. Vegetatively many Solanaceae - especially Solanoideae, the bulk of the family - are distinctive because the insertion of leaves, branches and flowers often does not seem to follow any regular sequence; paired leaves adjacent on the stem are common. The flowers often have porose anthers or the anthers as a group are connivent at the apex. The fruit is usually bicarpellate and is a septicidal capsule or a berry with flattened seeds; the calyx consists of connate sepals that persist and even enlarge in fruit.

Evolution. Stem group Solanaceae date from 66-65 million years before present, crown group (including Schizanthus and Duckeodendron) from 41-36 million years before present (Wikström et al. 2001); Janssens et al. (2009) date crown Solanaceae to 58±9.1 million years before present. A genome duplication event in Solanaceae has been dated from ca 50-52 million years before present (Schlueter et al. 2004).

New World Solanaceae are eaten by larvae of Nymphalidae-Ithomiinae butterflies (Ehrlich & Raven 1964; Drummond & Brown 1987; Willmott & Freitas 2006: Schizanthoideae, Goetzeoideae and Schwenkioideae may not be eaten); detailed coevolution seems not to be involved. Ithomiinae probably moved on to Solanaceae from Apocynaceae, and Solanum is especially important as a food source for caterpillars of this group (ca 70% records of neotropical Solanaceae food sources, ca 89% those of all Ithomiinae: Willmott & Freitas 2006; see also Brower et al. 2006). Some larvae may be distasteful because of the alkaloids, etc., of the leaves they eat, although Brown (1987) suggests that the noxious solanaceous chemicals guide oviposition and the feeding preferences of the larvae: "Though the butterflies may be able to recognise their food plants, biologists have greater difficulty in Solanaceae identification," (Brown 1987, p. 373). However, the adult butterflies are distasteful, but not because of these alkaloids, etc., rather because of the dihydroxypyrrolizidines that the adults obtain especially from Boraginaceae-Heliotropioideae and Asteraceae-Asteroideae (especially Eupatorieae); massive amounts (ca 20% dry weight) may be sequestered by the butterflies, which are initially quite palatable immediately after hatching (Brown 1987). The mimicry rings in which Ithomiinae are involved may be associated with particular solanaceous host plants (Willmott & Mallet 2004). Tobacco hornworm caterpillars prefer members of the [Solanoideae + Nicotianoideae] clade as food sources, although they didn't like Nicandra much; they died on Petunia, and didn't grow on Browallia and Brunfelsia. Other plant feeders show similar distinctive patterns (e.g. Fraenkel 1959), thus other sphingids are found here and on Oleaceae (Forbes 1958). Phytophagous Chrysomelidae beetles (perhaps especially Criocerinae) are notably more common on New World than Old World Solanaceae, perhaps suggesting an origin of the use of the family as a food source in the former area (Jolivet & Hawkeswood 1995; see also Hsiao 1986), while Chrysomelinae and Megalpodinae are also found on New World Solanaceae (Jolivet 1988). Criocerinae may have moved onto Solanaceae from monocots; the larvae are covered by faecal shields (Vencl & Morton 1999). In general, however, Solanaceae have multiple lines of defence and are avoided by most insect herbivores (Harborne 1986; Hsiao 1986).

Within Solanoideae, the Andean Iochrominae are notably diverse florally and have a variety of pollinators (Smith & Baum 2006). Zygomorphy and heteranthy (which in this context is really a kind of zygomorphy) have evolved several time in Solanum (Bohs et al. 2007); buzz pollination is common there (see Teppner 2005 for the pollination of the tomato; Harter et al. 2002; García et al. 2008) with over a million pollen grains being produced by a single flower (Anderson & Symon 1988); there is no nectar. Nierembergia (Petunioideae) has oil flowers (Coccucci 1991; see Tate et al. 2009 for a phylogeny). A number of taxa have quite large and complex stigmas (Cocucci 1991, 1995). Seed dispersal very much follows what fruit morphology might suggest (Knapp 2002b for a summary). The distinctively pungent capsaicanoids of chilis (Capsicum spp.) are involved in the protection of the fruit against the fruit-destroying Fusarium fungus (Tewskbury et al. 2008).

Petunia and Hyoscyamus, in different subfamilies, can be intergrafted (Taiz & Zeiger 2006).

Economic Importance. Chiles (Capsicum annuum were domesticated in Mexico, quite possibly in a number of places (Aguilar-Meléndez et al. 2009); other species of the genus are also economically important (Perry et al. 2007 and references); for the domestication of the tomato, see Bai and Lindhout (2007).

Chemistry, Morphology, etc. Lycium is recorded as accumulating glycine betaines, and some members at least are halophytic (Levin & Miller 2005). For alkaloids in Datureae, see Doncheva et al. (2006). Unusual stomata with degenerate guard cells seem often to have been reported in the family (Cammerloher 1920; D'Arcy & Keating 1973). Leaves in the fertile part of the stem of Solanaceae, perhaps especially in Solanoideae, are often geminate and/or branching is not simply axillary; Petunia can have ordinary-looking cymose inflorescences, but Schwenkia, Schizanthus and many other taxa have more or less recaulescent bracts, only one branch of the cymose inflorescence is developed at each node, or the two branches develop in different ways, etc., making interpretation of the construction of the plant very difficult (see especially Danert 1958, 1967; Child & Lester 1991 for a brief summary; Bell & Dines 1995). Goetzea has an odd growth pattern; its leaves are rather xeromorphic.

Physalis has notably inflated calyx surrounding the fruit in the development of which heterotopy of a foliar gene may be involved (He & Saedler 2007; cf. Hu & Saedler 2007); inflated calyces occur in some nine genera in total, although details of the pattern of evolution - and perhaps also loss - of this feature are unclear. In floral development, petal and stamen primordia together are lifted by zonal growth and the carpel primordia develop on a flat apex; in this respect there are some similarities between Solanaceae, Scrophulariaceae and Gesneriaceae, few with Montiniaceae (Huber 1980: 66-69; Ronse Decraene et al. 2000). For floral development in Datureae, see Yang et al. (2002). the flowers of Nolana have two long and two short stamens. The patterns of endothecial thickenings in the family are very diverse (Carrizo García 2002).

The two carpels so common in Solanaceae (but Nicandra has 3-5) are often in the plane of the first sepal initiated; this is one of the abaxial pair. Zygomorphy and heteranthy (which in this context is really a kind of zygomorphy) have evolved several time in Solanum (Bohs et al. 2007). The basic plane of symmetry in flowers like Salpiglossis and Schizanthus may be monocotyledonous, and the "abaxial" one or three stamens are sterile. Indeed, more or less well developed monosymmetry is quite common in Solanaceae, and is of the 3:2 sort (see Eichler 1875; Robyns 1931; Cocucci 1989; Knapp 2002a; Ampornpan & Armstrong 2002). Floral symmetry in both Convolvulaceae and Solanaceae needs study, especially given the suggestion by Ampornpan and Armstrong (2002) that in the latter the gynoecium is in the median plane, the parts being initiated slightly off the vertical plane of symmetry and the whole flower slightly rotated, the result being the odd petal is in the adaxial position. For a detailed study of the frequent loss of gametophytic incompatibility in Solanaceae, see Igic et al. (2006). Androgenesis, an uncommon condition in which the male gamete in maternal cytoplasm produces an embryo, has been recorded for at least one member of each of the three subfamilies Petunioideae, Nicotianoideae and Solanoideae - Petunia, Nicotiana, and Capsicum (Chat et al. 2003 for references).

Arabidopsis-type telomeres are absent from some Browallioideae (Sýkorová et al. 2003a). Cestreae in particular, which lack these telomeres, have chromosomes that at 7.21-11.51 µm long are considerably larger than those of the rest of the family, which are much smaller, e.g. 1.5-3.52 µm long in Nicotianoideae (Acosta et al. 2006; Tate et al. 2009); Cestreae also have n = 8 (Las Penas et al. 2006).

For general chemistry, see Hegnauer (1973, 1990, also 1966, 1990 as Nolanaceae), for hairs, esp. of Solanum, see Seithe (1962), for wood anatomy, see Carlquist (1987, 1988a) and Jansen and Smets (2001: vestured pits - do Petunioideae and Nicotianoideae have them?), for floral and inflorescence morphology, see Huber (1980), for branching patterns, see Danert (1958), for pollen morphology of zygomorphic taxa, see Stafford and Knapp (2006: not yet integrated with phylogeny), and of Hyoscyameae, see Zhang et al. (2009), for evolution of secondary metabolites, see Wink (2003 and references), for evolution of fruit and flower types, etc., see Knapp (2002) and Knapp et al. (2004), for calystegines (tropane alkaloids), see Dräger (2004), for the evolution of floral scent, see Martins et al. (2007), and for fruit anatomy, see Pabon Mora and Litt (2007).

Phylogeny. For early studies of relationships, see Olmstead & Palmer (1992) and Fay et al. (1998b). The grouping [Petunioideae [Solanoideae + Nicotianoideae]] is well supported in Olmstead et al. (1999), although less so in Olmstead and Santiago-Valentin (2003). However, in the summary tree of Olmstead and Bohs (2007), although there is a clade [Solanoideae + Nicotianoideae], immediately below it is a polychotomy including Petunioideae, Cestroideae and Schwenkioideae. Indeed, the relationships between the branches basal to this group have only weak support, Schwenkia may be sister to the rest of the family (Olmstead et al. 1999). However, using the nuclear gene SAMT (salicylic acid methyl transferase), Martins and Barkman (2005) found Schizanthus to be sister to the rest of the family, and with rather strong support, with Schwenkia weakly linked with Cestroideae (see also Olmstead & Bohs 2007). The Goetzioideae clade in the past has included Duckeodendron as sister to the rest, but with only moderate support (Santiago-Valentin & Olmstead 2001, 2003); here its relationships are unresolved. Furthermore, Wu et al. (2006) found a strongly supported grouping of [Solanoideae [Petunioideae + Nicotianoideae]], and although in this case no other clades of the family were included, Wu et al. (2006) noted that the sequences they included came from ten orthologous loci each on a different chromosome. (Wu et al. [2006] also dismissed the possibility that there had been a genome duplication either on the branch leading to Solanaceae or within Solanaceae themselves [cf. Blanc & Wolfe 2004].) The two-gene tree in a recent study by Olmstead et al. (2008) is rather like that of Martins and Barkman (2005), with relationships as follows: Schizanthoideae [Goetzioideae, Duckeodendron [[Cestroideae/Browallioideae, including Benthamiella et al.; their relationships have previously been unclear], Petunioideae, Schwenckioideae [Nicotianoideae + Solanoideae]]], but there is strong support for relationships between the last pair of taxa alone. The distinctive Sclerophylax is included in Solanoideae (Olmstead et al. 2008) - it has sessile flowers, an inferior oblique ovary with 2-3 ovules per carpel that are pendulous from the upper part of the loculus, and the calyx is accrescent, spinescent, and surrounds the 1-3-seeded fruit. Clearly there remain substantial uncertainties about relationships between major groupings within the family, and the account above is notably imperfect.

The limits of Solanum are to be expanded to include Cyphomandra and Lycopersicon; the hairs are often stellate and prickles are common (see Bohs 2005, 2007; Levin et al. 2006; Weese & Bohs 2007 [three genes, S. thelopodium sister ro rest, or unresolved in Bayesian analysis], Botany 2008: Botany without Borders 120-121. 2008; Poczai et al. 2008, for phylogenies). for relationships within the distinctive Nolana, see Tago-Nakawaza and Dillon (1999), Dillon et al. (2007) and Tu et al. (2008). Lycium may be paraphyletic, and then will need to be expanded to include Phrodus and Grabowskia (Levin & Miller 2005; Levin et al. 2007; Levin et al. 2009).

Classification. For generic descriptions and much else, see Hunziker (2001), although his suprageneric classification differs from that used here; Solanaceae Source includes information currently mostly about Solanum, but its coverage will expand. For the main outlines of the classification followed here, see Olmstead et al. (2008).

Previous Relationships. Huchinson (1973) placed Duckeodendraceae in Boraginaceae, but doubtfully; Cronquist (1981) kept it as a poorly-known family; Takhtajan (1997) placed it as a separate family in Solanales. Its carpels are oblique to the main axis of the flower (Kuhlmann 1934), as is appropriate for a genus in Solanaceae. Nolanaceae have often been separated from Solanaceae on account of their distinctive gynoecium; there are basically five carpels borne opposite the petals, but their number secondarily increases; the basic growth pattern is very like that of other Solanoideae (see also Eichler 1874).